I have, in this thesis, studied the interactions between gray-sided voles (Clethrionomys rufocanus) and tundra vegetation, on islands in, and mainland sites close to the lake Iešjávri, in northern Norway. As isolated islands are virtually free of predation, I have been able to compare plant-herbivore interactions in the presence and absence of predators. I transplanted vegetation from an island with predators and voles, to predator-free islands with and with out voles. The results reveal the existence of a terrestrial trophic cascade as voles had a severe impact on the transplanted vegetation on the predator-free islands, but only minor effects on the mainland where predators are present. Moreover, this study shows that plant defence was only a successful strategy when predators were present. Voles reduced the abundance of all available plants during winter on the predator-free islands. The results imply that cascading effects of predation are most important for well-defended plants with grazing-sensitive morphology as these plants escape herbivore impacts in the presence of predators but are vulnerable in their absence I studied the recovery of intensively grazed vegetation by building exclosures on islands that have been heavily grazed by voles for almost a decade.This study shows that the collective biomass of vascular plants recovered completely on three years, when voles were excluded. Although most species that are dominating the ungrazed vegetation recovered rapidly in the exclosures, the vegetation did not simply return to its ungrazed state. Herbaceous plants increased and there were pronounced differences in response among evergreen dwarf-shrub species. The semi-prostrate and tannin rich crowberry (Empetrum nigrum), showed the strongest recovery of all species, while the erect lingonberry (Vaccinium vitis-idaea) only showed weak signs of recovery. Thus, growth form determined the potential of plants to recover, whereas the trade-off between defensive investments and capacity to recover was weak or absent. I studied the interaction between gray-sided voles and their main winter food plant, bilberry (Vaccinium myrtillus) on islands in and mainland sites close to the lake Iešjávri. I compared the abundance, population structure and palatability of bilberry ramets between vole-free islands, islands with voles but no predators and mainland sites with both voles and predators. Voles decreased the abundance of bilberry on the mainland, but the effect was much stronger on predator-free islands. Bilberry was fairly tolerant to grazing as it partially compensated for the lost tissue by producing more new ramets. Moreover, a cafeteria experiment showed that voles preferred the ramets from predator-free islands, which is inconsistent with conjectures emphasizing inducible plant defenses. The vole-bilberry interaction lacks features of delayed density dependence that could explain the vole cycles. I conducted a clipping and fertilization experiment to further investigate the effects of herbivory on palatability of bilberry shoots. Fertilization decreased the concentration of condensed tannins in shoots of bilberry and voles preferred fertilized and clipped shoots. I found no indication of induced defense that could reduce the palatability of bilberry twigs in response to herbivory. The relationships between gray-sided vole densities, levels of invertebrate herbivory and chemical quality of leaves of Northern willow (Salix glauca) were studied on islands and mainland sites with contrasting vole densities. I found a positive correlation between level of invertebrate herbivory and vole density. The number of leaves per shoot, leaf size and leaf nitrogen content were also positively correlated with vole densities, while leaf C/N ratios were negatively correlated with vole densities. The positive correlation between vole densities and level of invertebrate herbivory is probably due to a facilitative effect of voles on invertebrate herbivores, mediated through changes in plant chemistry.

Forest management practices have changed the over-all structure of the Fennoscandian forest landscape resulting in a lack of suitable substrates for many wood-inhabiting species. The objectives of this thesis was to describe the colonization patterns of wood-inhabiting fungi, including the potential role of beetles as dispersal vectors, on different types of dead wood substrate and assess the importance of active measures in the forest landscape in order to restore biodiversity i.e. to increase the amount of dead wood and the use of restoration fire. The results clearly demonstrate the importance of restoration fire for wood-inhabiting fungi in a dry Pinus sylvestris forest. The general pattern for the majority of the species was a drastic decline the first two years after fire. However, after four years most of the species had recovered and were frequently found on logs strongly affected by the fire. The early fungal colonization patterns on fresh experimental Picea abies logs revealed no differences between managed forest stands and stands associated with nature reserves. After five years the species assemblage on the experimental logs was affected by stand age, forest site type, and distance to forest reserves. However, very few red-listed species colonized the logs in spite of being fairly common in the reserve stands. We conclude that the experimental period of only five years was too short to fully evaluate the possibilities to use experimental logs for threatened and red-listed species. We assessed the colonization patterns of different fungal functional groups based upon their different nutritional strategies namely mycorrhizal, saprotrophic on litter and humus, saprotrophic on wood causing white rot, and saprotrophic on wood causing brown rot. The results show that the fungal community undergoes a marked change in dominant nutritional strategies during the initial stage of the colonization process both after fire disturbance and on fresh un-colonized experimental logs. To which extent, saproxylic beetles are involved as passive or active vectors in the dispersal and colonization of wood-inhabiting fungi occurring on dead wood is poorly understood. The results clearly showed that some beetle species do discriminate between different fungal substrates and in particular, the bark beetle Dryocoetes autographus showed significant preference for wood with Fomitopsis rosea mycelium.

The aim of this thesis project was to elucidate patterns and processes associated with the biotic interactions in a natural plant–herbivore–parasitoid food web characterized by spatial and temporal heterogeneity with regard to species composition. The system examined is based on island populations of the perennial herb Meadowsweet (Filipendula ulmaria, Rosaceae), located in the Skeppsvik Archipelago. The area is subject to isostatic rebound, amounting to 0.85 cm per year; this makes it possible to calculate the age of the rising islands. Meadowsweet colonizes new islands when they are about 100 years old. Meadowsweet is consumed by two major herbivores in the study area: Galerucella tenella and Altica engstroemi (Coleoptera: Chrysomelidae). Both herbivores overwinter in the topsoil and successful colonization occurs when the islands reach a height that prevents the beetles from being removed or killed as a result of wave wash during the winter. I found that both herbivores significantly reduced individual plant fitness and population growth rate. A “cafeteria experiment” with Galerucella showed that this beetle discriminated between plants from different islands, avoiding plants from old islands which contained high concentrations of putative defence compounds, while readily accepting plants from younger islands which contained lower concentrations of these chemicals. Further, the plant species exhibited a trade-off between growth and production of the putative defence compounds. Taken together, these results were interpreted as providing evidence of herbivore-driven evolution of resistance in Meadowsweet. Further, laboratory studies suggested that Galerucella gradually includes a less preferred host plant (Rubus arcticus, Rosaceae) in its diet as Meadowsweet resistance increases. This implies that Galerucella drives its own host-breadth enlargement by selectively inducing a ‘rent rise’ in the original host, Meadowsweet. In a number of field studies I showed that the oligophagous parasitoid Asecodes mento (Hymenoptera: Eulophidae) has a strong positive effect on Meadowsweet seed set by removing large numbers of G. tenella larvae. This top-down effect is, however, altered by the presence of a close relative of G. tenella, namely G. calmariensis, which is monophagous on Purple loosestrife (Lythrum salicaria, Lythraceae). G. tenella experiences associational susceptibility when coexisting with G. calmariensis since the latter supports a higher and more fit pool of shared parasitoids and because Meadowsweet attracts a higher proportion of the shared parasitoid females than Purple loosestrife. This implies that G. tenella densities are very low in coexisting populations and that Meadowsweet experiences associational resistance and produces more seeds when co-occurring with Purple loosestrife. Thus, selection for increased resistance in Meadowsweet is likely to be relaxed in populations mixed with Purple loosestrife. I conclude that the evolution of plant resistance is likely to depend on the length of time and intensity of selection. When Meadowsweet colonizes new islands it experiences a period of enemy-free space; followed by a midlife and ageing with selection by herbivores. The intensity of this selection does, however, depend on the presence of additional plant and herbivore species.

A prey animal can use different strategies to avoid becoming eaten by predators. One such widely recognised strategy is the use of body colouration to decrease the risk of becoming detected, i.e. cryptic colouration. The principles of crypsis that I have studied are background matching, disruptive colouration and distractive markings. Further, I also studied the concealing effect of the visual background habitats. I used artificial prey items and backgrounds, and blue tits (Cyanistes caeruleus) as predators, to investigate prey concealment. In Paper I, I tested if high-contrast markings in prey coloration or in the background would result in a distracting effect. I found that such markings did increase prey search time, even when the prey markings were lighter or darker than the background. In Paper II, I studied the use of chromatic cues by predators when searching for prey. The birds easily detected prey that chromatically deviated from its background. Interestingly, background-matching prey was more difficult to detect when the colour scheme had low ultraviolet and high shortwave reflectance compared to when the reflectance bands were even. In Paper III, I studied optimisation of achromatic contrast within prey colour pattern and also the effect of shape diversity of background pattern elements on prey detection. I found that all prey types were more difficult to detect on the diverse background, but the level of contrast within prey pattern did not influence search times. In Paper IV, I further investigated how a prey should optimise its patterning with respect to background matching. I found that prey with repeated pattern elements was equally hard to detect as prey with more variable pattern. However, prey with a spatially regular pattern (aligned pattern elements) was easier to detect than prey with a spatially irregular pattern. In this paper I also found that high complexity of element shapes in the background, made the search task more difficult. / At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 1: Manuscript. Paper 2: Accepted

The oak district of Östergötland is one of the largest oak forests left in Europe and it harbours many threatened species dependent on old hollow trees. In this study, the saproxylic beetle Osmoderma eremita living in hollow oaks was used as a model species and parameterised for a metapopulation model (the incidence function model) at the level of individual trees. The aim was to set up a number of conservation management scenarios, where new oaks were added to increase the quality of the matrix and then simulate the resulting metapopulation occupancy, using three levels of dispersal. A real data set of hollow oaks was used as the base for modeling. Oaks were added to cells in a lattice covering the study landscape, where cell size, cell position and numbers of oaks per hectare were varied. The results show that a larger area of added oaks and a greater number of oaks per hectare will give a higher increase in total occupancy, no matter the dispersal, but the second smallest cell size used (3.4 km2 ) will give the highest response per effort for medium and high dispersal abilities, and smaller cells with greater number of oaks per area can be more effective. The spatial position of the increase in occupancy differs depending on dispersal ability of the focal species and should be considered in conservation management work. For unoccupied cells, the effort required to get the cell occupied will be smaller for cells having a greater number of oaks.

Studies have shown that air pollution, as well as bark fissure depth and sun exposure of a tree can have an effect on lichen growth as well as abundance of lichen species. The aim of this study was to find out the relative importance of these factors. 211 oaks in south eastern Sweden were surveyed for presence of 17 lichen species, as well as the total number of lichen species. Half of the trees were situated in urban areas and half in the countryside. For each tree the bark fissure depth was measured and the sun exposure of the trunk was estimated. The results showed a significant negative effect on a majority of lichens growing in an urban environment and a significant positive effect on a majority of lichens growing on trees with deep bark fissures. The sun exposure was a less important factor but could not be excluded as a factor affecting lichen occurrence. These results show the negative effect urbanization can have on rare lichen species, as well as be used in the development and maintenance for the nature reserves containing them

A tool for management and conservation of valuable forests in Sweden are WKH:s. In this study WKH:s different in size, connectivity, amount of dead wood and quality of logs were investigated for species richness of bryophytes and brackets on coniferous logs. The aim was to clarify which scales and features that was important for the diversity of species as well as for individual species. The results showed that the amount of dead wood was most important on the site scale, and some species were affected at the landscape scale, a positive effect of valuable tracts. On the scale of individual logs, factors such as diameter, sun exposure, succession stage, contact with ground and ground bryophytes cover was most important. Red-listed species preferred logs with large diameter and late successional stages. The conclusion was that the quality of the substrate and the amount of dead wood was most important, but the amount of WKH:s on the landscape scale was also important for some species.

Measuring and assessing vital rates such as births and deaths are prerequisites for understanding population dynamics. Vital rates may be affected by the density of individuals, even though the importance of density dependence on population dynamics has been debated for a long time. The mallard Anas platyrhynchos is one of the foremost game species in the Holarctic, with millions of birds in hunters’ bags annually. Still, basic knowledge about regulation of mallards’ vital rates is poor, and experimental studies on this topic are rare. In this thesis I have studied survival patterns and density dependence in mallards breeding in Sweden and Finland. Long-term ringing data from both countries were analysed for mortality patterns and causation, as well as for e.g. survival rate estimation. Most of the studies were, though, experiments run over two years involving manipulations of the density of nests, broods and/or adults, in southern and northern Sweden, comprising different biotic regions. Common response variables were survival of nests, ducklings and hens, mainly analysed with program MARK. About 90% of the recovered mallards in Finland and Sweden were hunting kills. However, survival rates were high, ranging from 0.66 to 0.81 for most groups (sex*age). The generality of density dependence was evident since such processes were detected in all studies. Consequently, depredation rate was higher in high nest density compared to low nest density. Survival of ducklings was density-dependent in both boreal and nemoral biotic regions, with food limitation being evident in the former region but not in the latter. In spite of their generality, density-dependent patterns varied within as well between years, and for nest predation rates also between landscape types. The findings about density dependence in breeding mallards in this thesis are novel since they are based on experiments. They are potentially of general interest for management because they embrace a variety of lakes in two geographically distant areas, each being representative for large temperate areas in the northern hemisphere. Detection of density dependence at the local scale may be important at larger scales, too, following the principle of ‘ideal preemptive distribution’ in a source-sink dynamic system.

Obligate brood parasites lay their eggs in nests of other species, relying on these host parents to care for their offspring. This phenomenon has been a curiosity amongst researchers since its first description and has become a model study system for testing such ideas as coevolution and species recognition. This thesis examines a few of the many questions that arise from this breeding system. The New Zealand Grey Warbler (Gerygone igata) and its brood parasite, the Shining Cuckoo (Chrysococcyx lucidus) are used as the main study species, although research on the eviction behaviour of Common Cuckoos (Cuculus canorus) has also been conducted. First, the current state of knowledge and recent discoveries regarding nestling rejection abilities of hosts is reviewed in chapter one. Second, a comparative study of New Zealand passerine begging calls has been conducted to test for begging call similarity between a brood parasite and its host, as well as developing a new technique for detecting the mode of coevolution that may be occurring in the parasite – host relationship. Parent-offspring communication in Grey Warblers is also examined to test for both parental and nestlings Parents use both alarm calls to warn offspring of potential danger, and also parental feeding calls to elicit a begging response from nestlings. By contrast, nestlings are able to signal both age and short term levels of need to parents through the acoustic structure of the begging call. The evolutionary costs and benefits of egg eviction behaviour in the Common Cuckoo are also tested. An experimental approach showed that egg eviction had a growth cost, but this cost was temporary and restricted to during and immediately after the egg eviction phase. A pattern of compensatory growth was observed after the eviction period, so that during the later nestling stages there was no difference in mass, and no difference in fledging age. Finally, variation in the Grey Warbler breeding biology and Shining Cuckoo parasitism rates are examined through both time and across latitudes. This research has shown a counterintuitive pattern of breeding phenology across latitudes. These patterns have implications for Shining Cuckoos both in terms of timing of available nests and host selection. Keywords: Begging call, breeding phenology, brood parasitism, coevolution, Common Cuckoo, eviction, Grey Warbler, parent-offspring communication, Shining Cuckoo.

Models of climate change predict that temperature will increase during the 21th century and the largest warming will take place at high northern latitudes. In addition to warming, predictions for northern Europe include increased annual precipitation and a higher proportion of the precipitation during winter falling as rain instead of snow. These changes will substantially alter the hydrology of rivers and streams and change the conditions for riverine communities. The warming is also expected to result in species adjusting their geographic ranges to stay within their climatic tolerances. Riparian zones and wetlands are areas where excess water determines the community composition. It is therefore likely that these systems will be highly responsive to alterations in precipitation and temperature patterns. In this thesis we have tested the predicted responses of riparian vegetation to climate-driven hydrologic change with a six year long transplant experiment (I). Turfs of vegetation were moved to a new elevation with shorter or longer flood durations. The results demonstrate that riparian species will respond to hydrologic changes, and that without rare events such as unusually large floods or droughts, full adjustment to the new hydrological regime may take at least 10 years. Moreover, we quantified potential effects of a changed hydrology on riparian plant species richness (II) and individual species responses (III) under different climate scenarios along the Vindel River in northern Sweden. Despite relatively small changes in hydrology, the results imply that many species will become less frequent than today, with stochastic extinctions along some reaches. Climate change may threaten riparian vegetation along some of the last pristine or near-natural river ecosystems in Europe. More extensive loss of species than predicted for the Vindel River is expected along rivers in the southern boreal zone, where snow-melt fed hydrographs are expected to be largely replaced by rain-fed ones. With a seed sowing experiment, we tested the differences in invasibility between open wetlands, forested wetlands and riparian zones (IV). All six species introduced were able to germinate and survive in all habitats and disturbance levels, indicating that the tested wetlands are generally invisible. Germination was highest in open wetlands and riparian zones. Increasing seed sowing density increased invasion success, but the disturbance treatments had little effect. The fact that seeds germinated and survived for 2 to 3 years in all wetland habitats indicates that wetland species with sufficiently high dispersal capacity and propagule pressure would be able to germinate and establish here in their respective wetland type. Our results clearly demonstrate that a changed climate will result in substantial changes to functioning, structure and diversity of boreal wetland and riparian ecosystems. To preserve species rich habitats still unaffected by dams and other human stressors, additional protection and management actions may have to be considered.